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Joe Paterno, who won more games than any other major-college football coach, and who became the face of Pennsylvania State University and a symbol of integrity in collegiate athletics only to be fired during the 2011 season amid a child sexual abuse scandal that reverberated throughout the nation, died Sunday in State College, Pa. He was 85.

His family announced his death in a statement released Sunday morning. The cause was lung cancer, according to Mount Nittany Medical Center, where he had been treated. Paterno’s family announced in mid-November that he had received a diagnosis of lung cancer after a visit to a physician regarding a bronchial illness a few days earlier. He lived in State College.

During his 46 years as head coach, as he paced the sideline in his thick tinted glasses, indifferent to fashion in his white athletic socks and rolled-up baggy khaki pants, Paterno seemed as much a part of the Penn State landscape as Mount Nittany, overlooking the central Pennsylvania campus known as Happy Valley.

When Penn State defeated Illinois, 10-7, on Oct. 29, 2011, the victory was Paterno’s 409th, and he surpassed Eddie Robinson of Grambling for most career victories among N.C.A.A. Division I coaches. Penn State’s president at the time, Graham B. Spanier, presented Paterno with a commemorative plaque in a postgame ceremony shown on the huge scoreboard at Beaver Stadium.

It would be Paterno’s last game. Within days his former defensive coordinator Jerry Sandusky was indicted and arrested on multiple charges of sexually abusing young boys extending back to his time on Paterno’s staff. On Nov. 9, Paterno and Spanier were fired by the university’s board of trustees because of their failure to go to the police after they were told of an accusation against Sandusky in 2002.

Paterno’s abrupt firing at 84 was something that could hardly have been imagined, although he had stubbornly clung to the spotlight at an age when most head coaches, whatever their renown, had retired.

He had held himself to an exceedingly high standard with what he called his “grand experiment”: fielding outstanding teams with disciplined players whose graduation rates far exceeded that at most football powers. His football program had never been tainted by a recruiting scandal. His statue stood outside Beaver Stadium alongside the legend “Educator, Coach, Humanitarian.”

Former players who succeeded in professional life far beyond the football field had told of their debt to him.

“Look how many go to medical school or law school,” said Bill Lenkaitis, a dentist in Foxborough, Mass., who played for Paterno in the 1960s and became a longtime center for the New England Patriots. “Look how many become heads of corporations.”

Paterno and his wife, Sue, were major benefactors of Penn State. During his nearly half-century as head coach, donors gave hundreds of millions of dollars to the university, helping to shape it into a major research institution, seemingly an outgrowth of his having made Penn State a national brand name through its football teams.

All-Americans by the Dozen

Paterno was a five-time national coach of the year. He had five unbeaten and untied teams, and he coached Penn State to the No. 1 ranking in 1982 and 1986. He took his Nittany Lions to 37 bowl games, winning 24 of them, and turned out dozens of first-team all-Americans.

On Saturday afternoons at Beaver Stadium, crowds exceeding 100,000 cheered on Nittany Lions players recruited by Paterno largely from Pennsylvania and nearby states. Many went on to stellar professional careers, among them running backs Franco Harris and Lydell Mitchell and linebacker Jack Ham.

Paterno had a career record of 409 victories, 136 defeats and 3 ties. He was surpassed only by John Gagliardi, who has won 484 games at Carroll College in Montana and St. John’s of Minnesota, coaching below the major-college level.

“Joe Paterno, to me, is maybe the greatest coach ever,” Bobby Bowden, the former Florida State coach, who is No. 3 in major-college coaching victories, told The Associated Press in 2006. “I mean that truly, and I said that 10 years ago.”

Penn State hired Bill O’Brien, the 42-year-old offensive coordinator for the New England Patriots, to replace Paterno this month. O’Brien has never been a head coach, and the decision was criticized by some former Penn State players.

The sexual abuse scandal that unfolded in November received national attention beyond the sports world and brought new attention to the issue of child predators and the obligation to care for their victims and to act quickly to prevent new assaults by serial offenders.

After the accusations against Sandusky surfaced, the university’s athletic director and the official who supervised the campus police were arrested on charges including perjury before a grand jury. They pleaded not guilty.

Sandusky had retired after the 1999 season, but he had retained access to the Penn State football facilities, where he escorted disadvantaged young boys being helped by a foundation he created. In what became the most notorious episode, a graduate assistant said he tried to explain to Paterno that he had seen Sandusky sexually abusing a boy in a team locker room late one night in March 2002. Paterno reported a version of the story to two superiors, saying that he set up a meeting between the graduate assistant and the athletic director and university vice president, but evidently never followed up.

Paterno’s firing, and his replacement by his assistant Tom Bradley on an interim basis, came hours after he said he would retire, although he planned to coach the final three games of the season after compiling an 8-1 record.

When word came of Paterno’s ouster, thousands of Penn State students rampaged through the downtown area near the university in support of him, engaging in sporadic vandalism and clashing with the police.

Paterno remained at his ranch house less than a mile from the campus on the Saturday after his firing, when Penn State played Nebraska at Beaver Stadium. Many students in the crowd of more than 100,000 came dressed as Paterno, with pants legs rolled up, white socks and thick-framed black eyeglasses. Hundreds wore T-shirts reading “Joe Knows Football.”

Rather than sprinting onto the field, the Penn State players walked from an end-zone tunnel hand in hand, and a moment of silence was dedicated to victims of child abuse. When Paterno’s son Jay, the quarterbacks coach, arrived at his parents’ home after the game — a 17-14 Nebraska victory — he pumped his fist in the air and chanted “We are. ...”

Joseph Vincent Paterno was born on Dec. 21, 1926, and grew up in the Flatbush section of Brooklyn. His father, Angelo, obtained a law degree after working as a court clerk and encouraged his children to pursue education.

Joe Paterno played football and basketball at Brooklyn Prep, a Jesuit-run high school, and studied the classics, which he said taught him the importance of being committed to an ideal.

He entered Brown University in 1946, played quarterback and defensive back, and majored in English. A younger brother, George, was a teammate, as he had been in high school.

Paterno planned to attend Boston University Law School upon graduation. But when Rip Engle, Brown’s head football coach, left to coach Penn State in 1950, Paterno went with him to coach the quarterbacks.

Paterno proved intense, brash and ambitious, and in June 1964 he was named Penn State’s associate football coach. He turned down an offer to become Yale’s head coach, then succeeded Engle when he retired in 1966.

Engle never had a losing season at Penn State, but although his teams appeared twice in the Liberty Bowl and twice in the Gator Bowl, they never made it to a high-profile bowl game. His last squad had a 5-5 record.

A Fast Ascension

Paterno was 5-5 as well in his first season as the head coach, but his team went 8-2-1 the next year, when he devised a new defensive system designed to confuse opposing offenses with shifts after the ball was snapped.

Then came the team’s breakout on the national football scene. Penn State went 10-0 in 1968 and in ’69 before victories in the Orange Bowl each season, and was unbeaten for 31 straight games before losing to Colorado in September 1970.

Early in 1973, Paterno turned down an offer to become the head coach of the New England Patriots. That June, he became the first Penn State football coach to deliver the university’s commencement address.

But Paterno was long stymied in his bid for a No. 1 ranking. His 1973 team, starring John Cappelletti in his Heisman Trophy season, was 12-0 without a top ranking. His 1978 squad was unbeaten in the regular season but lost to Alabama in the Sugar Bowl in the battle for the No. 1 spot. Paterno finally reached the top ranking with his 1982 and 1986 teams.

Paterno seemed indestructible, but in November 2006, he sustained a broken leg and a severe knee injury in a sideline collision during a game against Wisconsin. He had surgery and did not return to the sideline for the rest of the season. But he went onto the Beaver Stadium field for Penn State’s 2007 opener, beginning his 42nd season there as the head coach and surpassing the major-college longevity record at one program set by Amos Alonzo Stagg at the University of Chicago from 1892 to 1932.

Paterno was walking with a cane early in the 2008 season, in pain from leg and hip problems. He began coaching from the press box in October, then had hip-replacement surgery on Nov. 23. Penn State nonetheless made it to the Rose Bowl, with a No. 6 national ranking, and Paterno coached his team, again from a press box, in a loss to Southern California.

His 2009 team was 11-2 and beat Louisiana State in the Capital One Bowl, but his 2010 squad was 7-6 and lost in the Outback Bowl to Florida. Then came his final season, when Penn State had lost only once when the end came.

Usually preferring a conservative ground game, Paterno was best known as a strategist for his defensive units. Penn State was known as Linebacker U.

Paterno spent endless hours plotting strategy, and he was demanding at practice, where little escaped his notice. He was devoid of flourishes. His players wore high-top black shoes and white helmets with a simple blue stripe, forsaking fashionable school logos and little stars representing individual achievements, and their blue and white jerseys were unadorned by names on the back, all to project toughness and a selfless attitude.

“He’s a tremendous motivator,” said Kenny Jackson, who was a wide receiver on Paterno’s 1982 national championship team and who later coached under him.

In December 2007, Paterno became the third active coach inducted into the College Football Hall of Fame.

But as Paterno coached on into his 70s, his teams faltered. He had only one winning season from 2000 to 2004 and seemed overmatched in the Big Ten, which Penn State had joined in 1993 after years of success as an independent. There were several off-the-field incidents as his teams struggled — one quarterback needed stitches after a fight at an ice rink, and a center was held out of games for damaging an apartment wall by firing a bow and arrow. The episodes added to the air of disappointment surrounding Paterno, who prided himself on his athletes’ character.

Some critics also complained that Paterno’s run-oriented offense remained too conservative. High school stars were said to be looking elsewhere, assuming that Paterno would soon be gone.

In August 2004, Spanier, then the Penn State president, called Paterno “a brilliant man, the school’s greatest ambassador, the school’s best fund-raiser and a philanthropist.” (Paterno’s family had donated more than $4 million to the university, which named a wing of its expanded library for Paterno and his wife in 2000.)

But the Sunday before Thanksgiving Day 2004, Spanier and three top university administrators went to Paterno’s home, suggesting he retire, although he had been given a four-year contract extension the previous spring.

In an interview with The Pittsburgh Post-Gazette late in 2005, Paterno said he had told the university officials that Penn State was close to returning as a national power. “They didn’t quite understand where I was coming from or what it took to get a football program going,” Paterno said. “I said: ‘Relax. Get off my backside.’ ”

They did.

Paterno’s team lost only one game in 2005 and captured the Big Ten championship before defeating Florida State in the Orange Bowl in 2006.

“You guys think that you write something and all of a sudden I’m bothered?” Paterno told The Associated Press at the outset of the 2007 season. “I don’t read it.”

In addition to his wife and his son Joseph Jr., known as Jay, Paterno is survived by his sons David and Scott; his daughters, Diana Giegerich and Mary Kathryn Hort; and many grandchildren. George Paterno died in 2002. Joe Paterno’s wife and his five children were Penn State graduates.

The Classics as a Prism

For all his achievements, Paterno insisted that there was more to college football than results on the scoreboard. As he wrote in The New York Times on the eve of the 1989 season:

“A hard-fought, well-fought, hairline-close game is as classical in sports as tragedy in theater. A tragedy usually ends with the stage strewn with bodies from both sides of a struggle, and you can’t tell who won and who lost. Victory is contained within defeat, and defeat is contained within victory. That’s the way it is in the best of games. What counts in sports is not the victory but the magnificence of the struggle.”

Paterno said he was disappointed by the trustees’ decision to fire him after he announced he would retire at the end of the 2011 season. But in a statement just before the board acted, he expressed remorse over his personal failing. He chose a word that he had used in describing what he saw as the spirit of athletic competition back in 1989, but this time in a very different context — the episode that ended his career and left his reputation in tatters.

“This is a tragedy,” he said. “It is one of the great sorrows of my life. With the benefit of hindsight, I wish I had done more.”